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Geographic Variation in Hebe Macrantha (Plantaginaceae): Morphology and Flavonoid Chemistry

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Geographic Variation in Hebe Macrantha (Plantaginaceae): Morphology and Flavonoid Chemistry Tuhinga 15: 27–41 Copyright © Te Papa Museum of New Zealand (2004) Geographic variation in Hebe macrantha (Plantaginaceae): morphology and flavonoid chemistry Michael J. Bayly,1 Alison V. Kellow,1 Rebecca Ansell,1 Kevin A. Mitchell,2 and Kenneth R. Markham2 (1) Museum of New Zealand Te Papa Tongarewa, PO Box 467, Wellington, New Zealand [email protected] [email protected] (2) Industrial Research Limited, PO Box 31 310, Lower Hutt, New Zealand [email protected] [email protected] ABSTRACT: This paper investigates geographic variation in Hebe macrantha, endemic to mountains of South Island, New Zealand. It assesses the attributes, distribution, and appropriate taxonomic status of its two previously described varieties, the validity and circumscriptions of which have been questioned by some taxonomists. Morphometric analyses support the existence of two distinguishable entities that equate with previous circumscriptions of these varieties, whose continued recognition is recommended. Variety brachyphylla occurs in the north of the species’ geographic range, from the Anatoki Range, northwest Nelson, to the Hanmer Range, north Canterbury. Variety macrantha is geo- graphically more widespread and morphologically more variable. It occurs in the south of the species’ range, south and west from Mt Haast (southwest Nelson) and Lewis Pass (south Nelson/north Canterbury), as well as at Lake Tennyson (southeast Nelson), the only locality at which specimens assigned to both varieties occur. The two varieties are morphologically most similar at localities close to their geographic interface. Patterns of variation in leaf flavonoids neither strongly support nor contradict recognition of the two varieties. Analysed samples share a similar and, within Hebe, distinctive flavonoid profile, and the distribution of two flavonoid glycosides is partly correlated with the morpholog- ical circumscription of varieties (the correlation is incomplete in samples taken near the geographic interface of varieties). A distribution map and a table of key morphological differences between varieties are provided. KEYWORDS: Hebe macrantha, Scrophulariaceae Plantaginaceae, flavonoids, New Zealand flora. 28 Tuhinga, Number 15 (2004) Fig. 1 Some morphological features of Hebe macrantha: (A) habit of a plant of var. brachyphylla on Mt Arthur; (B) sprigs of var. macran- tha (left) and var. brachyphylla; (C) top view of shoot apex, showing that young leaves diverge early (i.e. there is no prominent ‘lead bud’ like that seen in many hebes); (D) lower and upper leaf surfaces of var. macrantha (top) and var. brachyphylla; (E) inflorescence showing frontal view of flower; (F) lateral view of flower; (G) inflorescence (left) and infructescence; (H) septicidal (left) and loculicidal views of capsule (voucher specimens: A, from same population as WELT 80740; B (left), D (upper), WELT 82429; B (right), E, F, G, WELT 83544; C, D (lower), WELT 82554; H, WELT 83545). Variation in Hebe macrantha 29 Introduction Kellow et al. 2003a, b). It aims to identify and describe patterns of geographic variation within H. macrantha Hebe macrantha (Hook.f.) Cockayne et Allan is a distinc- using data on morphology and leaf flavonoids, and to re- tive sub-shrub from alpine regions of South Island, New assess its infraspecific classification. In particular, it seeks Zealand. It is readily distinguishable from other members to determine whether or not there are grounds for the of Hebe by its much larger flowers (see Figs. 1E and 1F), recognition of two taxa, and if so, to identify clearly their fruits that are laterally compressed (with the septum at distinguishing features, map their distributions, and assess right angles to the widest diameter; see Fig. 1H), and leaves their appropriate taxonomic rank. that are prominently petiolate (see Fig. 1D), have toothed This study was initiated primarily because an initial margins, and diverge early in bud (see Fig. 1C). Moore (in survey of specimens suggested that the leaf dimensions Allan 1961), in an informal infrageneric classification of specified by Moore (in Allan 1961) were insufficient to Hebe, placed H. macrantha in the monotypic grouping identify reliably all specimens to variety rank. A further “Grandiflorae”, this name referring to the characteristically motivation was the opinion of Prof. P. Garnock-Jones large flowers. Alternatively, Heads (1987), considered H. (Victoria University of Wellington), based primarily on macrantha best removed from Hebe and placed in field observations of morphological variation in the area Parahebe, whose members it resembles in many morpho- around Lake Tennyson, south Nelson Province (personal logical features. More recent analyses of nuclear ribosomal communication 1997), that there were probably insuffi- DNA sequences (Wagstaff & Garnock-Jones 1998, cient grounds for recognition of two taxa, and that the Wagstaff et al. 2002) group H. macrantha with other classification of H. macrantha warranted thorough assess- members of Hebe, sister to all other species of the genus ment. Heads (1994a) also suggested that ‘the two varieties (as defined by Wagstaff et al. 2002, and followed here). of the species and their distribution require revision’. Cheeseman (1906) reported that specimens of Hebe macrantha (as Veronica macrantha) ‘from Mount Arthur and other parts of the Nelson District have shorter, broad- Materials and methods er leaves, more numerous racemes and smaller flowers than Morphometric analysis is usual in Canterbury and Otago …’, and provided the variety name brachyphylla to distinguish these collections Fifty-one herbarium specimens from WELT and CHR from more typical forms. Cockayne and Allan (1926) (see Appendix 1; herbarium abbreviations follow agreed with the presence of two varieties of H. macrantha Holmgren et al. (1990)), covering most of the geographic (using the name var. vera for the typical form), and added range of Hebe macrantha, were scored for 16 morphologi- that var. brachyphylla ‘occurs not only in the North-western cal characters (see Table 1). The geographic distribution of Botanical District, as given in the Manual, but also in the these specimens is shown in Fig. 4, although three speci- wetter part of the North-eastern District’. Moore (in Allan mens – WELT 13110 (‘western Amuri District’, presum- 1961) also recognised two varieties in Hebe macrantha, and ably somewhere to the west of Hanmer Springs), WELT further defined them, giving leaf dimensions of c. 1.5–2.5 13101 (‘Ashburton mountains’), and WELT 13111 x 0.5–1.0 cm for var. macrantha and c. 1.3–1.5 x 0.9– (‘Otago’) – have only vague locality information and could 1.0 cm for var. brachyphylla. This classification, including not be mapped. Unless each of the fragments on an two varieties, has been followed in subsequent, general herbarium specimen was known to be from a single indi- works on the New Zealand flora (e.g. Eagle 1982, Druce vidual (e.g. in the case of MJB collections), or fragments 1993, Wilson & Galloway 1993, Mark & Adams 1995). were very small or were uniform in appearance, only one Distribution maps for the two varieties are given by Heads fragment was scored per specimen. In one case, WELT (1994a) and Macdonald (1982; with the two maps labelled 82420, two fragments on the one sheet were known to be the wrong way around), with the former providing no data from separate plants, and each was treated as a separate on morphological circumscription, and the latter largely specimen (specimens 12a and 12b in Appendix 1). repeating details given by Moore (in Allan 1961). Vegetative characters were measured for each of 10 leaves The study presented here is part of ongoing work and 10 internodes per specimen. Characters of flowers and toward a revised classification of Hebe (e.g. Bayly et al. inflorescences were measured for up to eight flowers or 2000, 2001, 2002, 2003; Garnock-Jones et al. 2000; inflorescences per specimen (where available). Character 30 Tuhinga, Number 15 (2004) Table 1 Characters used in the morphometric analysis. to the Kolmogorov-Smirnov test for normality (Sokal & Rohlf 1969) and Levene’s median test for homogeneity of variance (Levene 1960). For characters that passed tests for Internode length (INT) (mm) normality and equal variance (P > 0.05), t-tests were used Lamina length (LL) (mm) to compare groups. For characters that failed either test, the Lamina width (LW) (mm) Mann-Whitney Rank Sum Test was used to compare Lamina length/lamina width (LL/LW) groups. These analyses were performed using SigmaStat for Lamina length/total leaf length (LL/TLL) Windows Version 2.0. Distance from leaf base (including petiole) to widest point (DWP) (mm) Examination of additional Distance to widest point/total leaf length herbarium specimens (including petiole) (DWP/TLL) Results of the morphometric analysis were assessed by Petiole length (PET) (mm) reference to further herbarium material. All specimens of Number of teeth on one side of leaf (the bigger Hebe macrantha at WELT and CHR (95 additional speci- number if uneven) mens) were assessed for the five most discriminating mor- Peduncle length (mm) phological characters, and on the basis of their geographic Rachis length (mm) origin, to determine whether variation among all speci- Rachis length/total inflorescence length mens was congruent with, and adequately reflected by, Number of flowers per inflorescence that shown in the morphometric study. Length of lowermost bracts on inflorescences (mm) Pedicel length (mm) Analysis of leaf flavonoids Calyx
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